Pharaxonotha Reitter, 1875

Skelley, Paul E., Tang, William & Pérez-Farrera, Miguel Angel, 2022, Review of Pharaxonotha Reitter (Coleoptera: Erotylidae: Pharaxonothinae) inhabiting the cycad genus Dioon Lindl. (Cycadales), with descriptions of nine new species and comments on P. kirschii Reitter, Insecta Mundi 2022 (917), pp. 1-41 : 4-7

Key to species of Pharaxonotha Reitter on Dioon

1. Head in lateral view robust, thick, prominent ventrally, convex dorsally ( Fig. 1A ); post-ocular ridge present (temple), small, tooth-like in dorsal view ( Fig. 1A , 2A ); male with small tooth or denticle at apex of last abdominal ventrite ( Fig. 2D ); body strongly glossy, robust, dark red-brown to black; southern US to Central America, occasionally associated with cycads [kirschii group].................. 2

— Head in lateral view narrowed, more flattened anteriorly, not prominent ventrally, flattened dorsally and ventrally ( Fig. 1B–D ); post-ocular ridge indistinct, mostly lacking ( Fig. 2B–C ); male lacking modified last abdominal ventrite; body weakly glossy to distinctly dulled, flattened, red-brown to yellow-brown, rarely dark brown to black; widespread in the Neotropics, associated with cycad cones............................................................................... 4

2(1). Antennomere XI visibly larger, more robust than the small antennomere IX ( Fig. 2E ); body of most dark-brown to nearly black; male with distinct denticle on last abdominal ventrite; USA (Texas, Louisiana) to Panama in nature, Europe and elsewhere in plant products, free living................................................................................... P. kirschii Reitter

— Antennomere XI visible smaller than the robust antennomere IX ( Fig. 2F ); body of most red-brown, rarely black; male with indistinct denticle on last abdominal ventrite; Mexico, associated with cycads.............................................................................. 3

3(2). On female, fuscous spot on elytra covering <½ area of elytra ( Fig. 11A ); spermatheca apex gradually curving to a rounded point; male paramere width in lateral view> 3.4× own length ( Fig. 11I ); inhabiting the Dioon angustifolium Miq. complex in Tamaulipas, Mexico.... P. sclerotiza , new species

— On female, fuscous spot on elytra covering> ¾ area of elytra ( Fig. 10A ); spermatheca apex abruptly curving to a truncated point; male paramere width in lateral view <3.4× own length ( Fig. 10J ); inhabiting Dioon edule Lindl. in Veracruz, Mexico.................. P. dimorpha , new species

4(1). Dorsally, hairs emerging from punctures visible under low magnification (<30×), procumbent, length nearly equaling or exceeding distance to nearest puncture; inhabiting Dioon spp. in Mexico.... 5

— Dorsally, body appearing glabrous under low magnification (<30×) or, when visible, length of hairs associated with punctures distinctly less than distance to nearest puncture; inhabiting Zamia spp. , Ceratozamia spp. , and Microcycas sp. from Florida and the Caribbean, and Mexico south to Bolivia............................................... Go to couplet 2 in Skelley and Tang (2020)

5(4). Eyes small, finely facetted, 3–4 facets = length of antennomere II ( Fig. 1B ); eyes widely separated, head width/dorsal interocular distance (HW/DI) = 1.33–1.44 (1.54); head width/ventral interocular distance (HW/VI) <1.73; anterior pronotal angles broadly rounded, not projecting forward beyond anterior margin; body alutaceous, strongly dulled, some populations bicolored with elytra black to dark brown and contrasting with orange-brown pronotum; inhabiting Dioon califanoi De Luca & Sabato , Dioon caputoi De Luca & Sabato , Dioon purpusii Rose in northern Oaxaca and Puebla [bicolor group]................................................... P. bicolor , new species

— Eyes larger, encroaching on frons and gular region, coarsely facetted, 2–3 facets = length of antennomere II ( Fig. 1C–D ); eyes more narrowly separated, HW/DI = 1.48–1.72; HW/VI> 1.92; anterior pronotal angles sharply rounded, projecting forward beyond anterior margin; body weakly dulled, uniformly orange-brown to yellow-brown or pronotum slightly darker than elytra...................... 6

6(5). Body large, length> 4.7 mm; pronotal punctures large, impressed, sharply defined ( Fig. 6D ); inhabiting Dioon stevensonii Nic. -Mor. & Vovides in Guerrero, Mexico [occidentalis group, in part]........................................................................ P. gigantea , new species

— Body smaller, length <4.5 mm; pronotal punctures small, not sharply defined; inhabiting other species of Dioon in other Mexican states....................................................... 7

7(6). Head more flattened between eyes, in lateral view hardly visible above and below eyes ( Fig. 1C ) ; transverse occipital line [vertexal line] absent, with row of punctures that may appear as line ( Fig. 2B ); male protarsus and metafemur distinctly more swollen than female; male meso- and metatibia median margin slightly concave with small inwardly directed tooth at apex; inhabiting Pacific drainage in Jalisco, Oaxaca and Chiapas or in northeastern Oaxaca [occidentalis group, part]....... 8

— Head convex between eyes, in lateral view readily visible above and below eyes ( Fig. 1D ) ; transverse occipital line [vertexal line] present ( Fig. 2C ); male protarsus and metafemur weakly swollen compared with female; male meso- and metatibia median margin straight to weakly convex and lacking tooth at apex; inhabiting Atlantic drainage in Hidalgo, Nuevo Leon, Querétaro, San Luis Potosí, Tamaulipas and northern Veracruz [vovidesi group]..................................... 10

8(7). Posterior lateral margin of eye with small distinct fringe of setae separated from facets ( Fig. 2B ), best visible from dorsal profile; pronotal length/elytral length (PL/EL) = 0.27–0.33; male tegmen with parameres in dorsal view with pronounced bulge on outer margin; inhabiting Dioon holmgrenii De Luca, Sabato & Vázq. Torres , Dioon merolae De Luca, Sabato & Vázq. Torres , Dioon tomasellii De Luca, Sabato & Vázq. Torres in Chiapas, Jalisco and western and southern Oaxaca ............ 9

— Posterior lateral margin of eye without distinct fringe of setae; (PL/EL) = 0.32–0.37; male tegmen with parameres in dorsal view with margins nearly straight; inhabiting Dioon spinulosum Dyer ex Eichler in northeastern Oaxaca .......................................... P. woodruffi , new species

9(8). Elytral setae fine, short, reaching next puncture in length ( Fig. 8A–B ); HW/DI = 1.54–1.67, HW/VI = 1.92–2.44; inhabiting Dioon holmgrenii and Dioon merolae in western Oaxaca and Chiapas .................................................................... P. occidentalis , new species

— Elytral setae prominent, long, surpassing next puncture in length ( Fig. 7A–B ); HW/DI = 1.69–1.72, HW/ VI = 2.64–2.82; inhabiting Dioon tomasellii in Jalisco ................... P. novoai , new species

10(7). Head with transverse occipital line [vertexal line] long laterally in most, most with line surpassing half distance between stridulatory file and eye, often curving anteriorly near eye; mean pronotal length/ width (PL/PW) = 0.69–0.70 (multiple specimens required); in females spermatheca elongate, ratio of spermatheca length to width at basal third> 4.5, base not abruptly swollen (see Fig. 2G ); inhabiting the Dioon angustifolium complex north of the Mexican Transvolcanic Belt in Hidalgo, Nuevo Leon, Querétaro, San Luis Potosí and Tamaulipas .................... P. fawcettae , new species

— Head with transverse occipital line [vertexal line] short laterally in most, most with line stopping about half distance between stridulatory file and eye, not curving anteriorly; mean PL/PW = 0.71–0.74 (multiple specimens required); in females spermatheca short and compact, ratio of spermatheca length to width at basal third <3.5, base abruptly swollen, swelling often quadrate-shaped; inhabiting Dioon edule south of the Mexican Transvolcanic Belt in Veracruz .... P. vovidesi , new species

Figures

Figure 1. Pharaxonotha heads, lateral view. A) P. kirschii, Mpio. Jalpan, Querétaro, Mexico. B) P. bicolor, paratype, San Andres Pápalo, Oaxaca, Mexico. C) P. occidentalis, paratype, El Campanario, Chiapas, Mexico. D) P. vovidesi, paratype, Mt. Oscuro, Veracruz, Mexico.

Figure 2. Pharaxonotha body parts. A–C) Dorsal view of head. A) P. kirschii, Mpio. Jalpan, Querétaro, Mexico. Arrow points to temple. B) P. occidentalis, paratype, El Campanario, Chiapas, Mexico. C) P. vovidesi, paratype, Mt. Oscuro, Veracruz, Mexico.D) P. kirschii, abdominal ventrites, Mpio. Jalpan, Querétaro, Mexico. Arrow points to tooth on ventrite 5. E–F) Antennal clubs. E) P. kirschii, Santa Isabel, Chiapas, Mexico.F) P. sclerotiza, holotype. G) Spermatheca of Pharaxonotha fawcettae from Jalpan de Serra, Querétaro, Mexico indicating the diagnostic structures and orientations used in descriptions.

Figure 6. Pharaxonotha gigantea from Achotla, Guerrero, Mexico (male holotype). A–C) Dorsal, lateral and ventral habitus. D–E) Head, dorsal and ventral views. F) Tegmen attached to median lobe, ventral view top and lateral view bottom. G) Parameres of tegmen, ventral view top and lateral view bottom. H) Genital capsule, ventral view.

Figure 7. Pharaxonotha novoai from Cobo Corrientes, Jalisco, Mexico (A–I: male holotype; J–K: female allotype). A–C) Dorsal, lateral and ventral habitus. D–F) Head, dorsal, lateral and ventral views. G) Genital capsule attached to tegmen and median lobe, ventral view. H) Tegmen attached to median lobe, lateral view. I) Parameres of tegmen, ventral view top and lateral view bottom. J) Spermatheca with retracted ovipositor, ventral view. K)

Figure 8. Pharaxonotha occidentalis from El Campanario, Chiapas, Mexico (A–D, F: female allotype). A–C) Dorsal, lateral and ventral habitus. D–F) Head, dorsal, lateral and ventral views. G) Tegmen attached to median lobe, ventral view top and lateral view bottom. H) Parameres of tegmen, ventral view top and lateral view bottom. I) Retracted ovipositor, ventral view. J) Spermatheca (partially inflated).

Figure 10. Pharaxonotha dimorpha from El Farallón, Veracruz, Mexico (B–G: male holotype). A) Female allotype, dorsal habitus. B–D) Male dorsal, lateral and ventral habitus. E–F) Male head, dorsal and ventral views. G) Tegmen attached to median lobe and genital capsule, lateral view. H) Male genital capsule, dorsal view.I) Tegmen attached to median lobe, lateral view top and ventral view bottom. J) Parameres of tegmen, ventral view top and lateral view bottom. K) Retracted ovipositor, ventral view. L) Spermatheca (fully inflated).

Figure 11. Pharaxonotha sclerotiza from Gómez Farias, San Luis Potosí, Mexico (B–I: male holotype). A) Female allotype, dorsal habitus. B–D) Male dorsal, lateral and ventral habitus. E–F) Male head, dorsal and ventral views. G) Tegmen attached to median lobe and genital capsule, ventral views. H) Tegmen attached to median lobe, lateral view. I) Parameres of tegmen, ventral view top and lateral view bottom. J) Spermatheca and retracted ovipositor, ventral view. K) Spermatheca (fully inflated).